The Botrychiopsis Genus and Its Biostratigraphic Implications in Southern Paraná Basin

Anais da Academia Brasileira de Ciências (2003) 75(4): 513-535 (Annals of the Brazilian Academy of Sciences) ISSN 0001-3765 www.scielo.br/aabc

The Botrychiopsis genus and its biostratigraphic implications in Southern Paraná Basin

ANDRÉ JASPER1, MARGOT GUERRA-SOMMER2, MIRIAM CAZZULO-KLEPZIG2 and RUALDO MENEGAT2 1Setor de Botânica e Paleobotânica, Museu de Ciências Naturais, Centro Universitário UNIVATES (SBP/MCN/UNIVATES),95900-000 Lajeado, RS, Brasil 2Instituto de Geociências, Universidade Federal do Rio Grande do Sul (IG/UFRGS) 91501-970 Porto Alegre, RS, Brasil

Manuscript received on February 3, 2003; accepted for publication on July 23, 2003; presented by Alcides N. Sial

ABSTRACT Botrychiopsis has been considered an important floristic element of Westphalian/Artinskian associations of the Paraná Basin. The occurrence of Botrychiopsis in roof-shales of the Rio Bonito Formation in Southern Paraná Basin (Quitéria area), supported by the identification of Botrychiopsis valida, enlarges the genus biochron. Consequently, the stratigraphic hierarchy for Botrychiopsis plantiana and Botrychiopsis valida was defined for the Paraná Basin. Although it is climatically controlled and related to a deglaciation icehouse stage, stratigraphic distribution of the genus presents a substantial climate tolerance, from cold/cool to warm/temperate conditions. A new phytostratigraphic zonation is proposed for the southern portion of the basin that includes the Botrychiopsis Zone (Asselian/Kungurian), which is subdivided into the Botrychiopsis plantiana (Asselian/Artinskian) and Botrychiopsis valida (Late Artinskian/Kungurian) subzones. Key words: Botrychiopsis, biostratigraphy, Paraná Basin, Permian, palaeoclimatology, Gondwana.

INTRODUCTION sic – Early Cretaceous and Late Cretaceous) encom- pass continental strata and volcanic rocks. Each The intracratonic Paraná Basin covers 1.700.000 megasequence corresponds to previously proposed square kilometers of East and Central South lithostratigraphic units. America (1.100.000 square kilometers in Brazil) and The Carboniferous – Early Triassic (CET) encloses Paleozoic, Mesozoic and, locally, Ceno- megasequence represents a second-order transgres- zoic sedimentary and volcanic rocks. According to sive – regressive cycle. It contains a basal transgres- Milani et al. (1998), the Paraná Basin comprises sive unit overlain by a regressive succession. The six stratigraphic megasequences bounded by interre- basal succession includes the Itararé Group and Rio gional unconformities (sensu Vail et al. 1977). The Bonito and Palermo formations. three lower megasequences (Ordovician – Silurian, Botrychiopsis plantiana, recoverd from thick Devonian and Carboniferous – Early Triassic) con- sandstone and siltstone strata, was recorded in the sist of transgressive – regressive cycles whereas the megafloras of the lower part of the Brazilian Gond- three upper megasequences (Late Triassic, Juras- wana succession, which comprises lowland glacio- Correspondence to: André Jasper continental deposits of the Itararé Group (Millan E-mail: [email protected]

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1975, 1979, Cazzulo-Klepzig and Guerra-Sommer shales in Southern Paraná Basin (Quitéria area), as 1983, Zampirolli 2001). discussed herein, allow to infer the relationship be- The synonymization of distinct taxa to this tween the biostratigraphic distribution of this genus morphogenus, which is characterized by heteromor- and its tolerance to climate changes. In this case, cli- phy, took place for more than a century (1844-1971) mate change has been associated with a deglaciation during which several Earth Science paradigms were phase of an icehouse period and is represented by an established and abandoned, from the Fixist models evolution from cool-temperate to warm-temperate to Plate Tectonics. The main studies that produced climate. The validation of this hypothesis would the current genus conception are presented in Fig. 1. then broaden the current parameters used to inter- Studies of Archangelsky and Arrondo (1971) pret the climatic conditions favorable for the devel- demonstrated the biogeographic and biostrati- opment of these plants. graphic importance of the genus in Gondwana suc- This paper comprises the first step of a larger cessions. These sequences occur in Argentina project that aims to evaluate the chronostratigraphic (Sessarego and Césari 1986, Archangelsky et al. significance of the Botrychiopsis within Gondwana. 1987, Archangelsky and Cúneo 1987, Andreis and It was carried out to verify earlier taxonomic and Archangelsky 1996); Brazil (Millan 1975, 1979, biostratigraphic descriptions of Botrychiopsis forms 1987a,b, Rösler 1978, Guerra-Sommer and Caz- of the Southern Paraná Basin. This revision was re- zulo-Klepzig 1981, Guerra-Sommer and Cazzulo- quired due to the reduced nature of previous descrip- Klepzig 1993); South Africa (Rayner 1985, 1986, tions, which did not define the diagnostic differences Rayner and Coventry 1985, Anderson and Ander- between previously identified and original material. son 1985, Kovács-Endrödy 1991), India (Srivastava 1997) and Australia (Rigby 1973, 1993, Retallack REVIEW OF THE Botrychiopsis OCCURENCE 1980). IN SOUTHERN PARANÁ BASIN The paleofloristic assemblages show a homo- Forms related to Botrychiopsis have been identified geneous composition dominated by foliar organs of in Rio Grande do Sul State in Faxinal, Base of Morro plants with arborescent habit as well as remnant Papaléo and Quitéria outcrops (Fig. 2). shrub-like plants, such as Botrychiopsis plantiana. The Faxinal outcrop, described by Morgenthal Shrub-like plants identified as Botrychiopsis et al. (1970) and later by Andreis et al. (1979), is plantiana have also been recorded in the southern- located in the northeastern portion of the Barão do most portion of the Paraná Basin, within the Glos- Triunfo topographic Sheet, close to the confluence sopteris Flora and are associated with fluvial, delta of the Grande and Faxinal creeks, about 10 km west- and estuarine sedimentary rocks at the base of the northwestward from the Mariana Pimentel District. Rio Bonito Formation (Pasqualini et al. 1986). The exposure includes variable amounts of The presence of Botrychiopsis in Paleozoic sandstone and mudstone related to the uppermost Gondwana floras of the Paraná Basin has usually part of the Itararé Group in the Rio Grande do Sul been associated with tundra and taiga environments State. It represents the westernmost exposure of a (Archangelsky 1971, 1978, 1984, Rocha-Campos paleovalley that extends from Potreiro Grande, 4 kM and Archangelsky 1985, Retallack 1980, 1999). eastwards from the Mariana Pimentel District. In their description of the Glossopteris Flora Andreis et al. (1979) recognized two infor- Early Permian evolution in Southern Brazil, Guerra- mal siltstone facies (white and grayish brown silt- Sommer et al. (1991, 2001) regarded Botrychiop- stone facies). Isolated Rubidgea-type leaves, Botry- sis plantiana as a remnant plant from a rigorous chiopsis fronds and small platispermic seeds rep- periglacial cold climate. resent most of the quantitatively poor megafloristic The first records of Botrychiopsis within roof- association. Minor amounts of the Cordaites, Glos-

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Taxa Quotation Author Year Botrychiopsis weissiana Kurtz 1895 Botrychiopsis weissiana Kurtz 1921 Adiantites (?) robustus Wallkom 1934 Cardiopteris cf. frondosa Wallkom 1934 (?) Botrychiopsis weissiana ? Noeggerathia sp. Wallkom 1934 (?) Botrychiopsis weissiana Kurtz & Frengüelli 1944 Gondwanidium plantianum Gerth & Frengüelli 1944 ? Adiantites sp. Frengüelli 1946 Gondwanidium plantianum Gerth & Frengüelli 1946 Botrychiopsis weissiana Archangelsky & Arrondo 1971 Odontopteris plantiana Carruthers 1869 Neuropteridium validum Feistmantel & Kurtz 1895 Neuropteridium validum Feistmantel & Seward 1903 Neuropteridium validum Feistmantel & Arber 1905 Neuropteridium plantianum White 1908 Botrychiopsis plantiana Neuropteridium plantianum Lundqvist 1919 Neuropteridium validum Feistmantel & Kurtz 1921 Neuropteridium validum Kurtz 1921 Neuropteridium plantianum Dolianiti 1953 Botrychiopsis plantiana Archangelsky & Arrondo 1971 Gondwanidium plantianum Rigby 1973 Otopteris ovata Mc’Coy 1847 Neuropteris valida Feistmantel 1876 Neuropteris valida Feistmantel 1879 Neuropteridium validum Feistmantel 1880 Botrychiopsis valida Godwanidium validum Gothan 1927 Gondwanidium validum Gothan 1941 Gondwanidium validum Gothan 1966 Botrychiopsis valida Archangelsky & Arrondo 1971 Botrychiopsis ovata Gould 1976 Botrychiopsis ovata Rettalack 1980

Fig. 1 – Current Botrychiopsis genus conception.

sopteris, Gangamopteris and Ginkgoites genera as iments, deposited during the deglaciation produced well as articulate stems and conifer branches are also by warming of a glacial climate. present. The specimens hereby presented were previ- According to Andreis et al. (1979) both white ously studied by Guerra-Sommer et al. (1980) and and grayish brown facies represent the final filling Cazzulo-Klepzig and Guerra-Sommer (1983) and stage of large lakes, and considered then as related are stored (samples PB 3097 e PB 3098) in the Pale- to the end of the Itararé Group sedimentary cycle obotanic Sector of the Earth Sciences Institute of the (Corrêa da Silva 1970 and Bossi and Piccoli 1979). Federal University of the Rio Grande do Sul State According to Milani et al. (1998), these sedi- (UFRGS). ments correspond to lowland glaciocontinental sed- Samples from Faxinal include basal fragments

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Fig. 2 – Localization of the Faxinal (1), Base of Morro Papaléo (2) and Quitéria (3) outcrops. of long bipinnate fronds (3,9 to 8,2 cm long and nate their bipinnate aspect, with pines ranging from 0,6 to 2,9 cm wide). They present a main robust oblong and rhomboidal at their base (Fig. 3e and f) to rachis, between 0,6 cm (proximal part) and 0,2 cm oblong-elliptical or elliptical at their apex (Fig. 3a), (distal portion) wide, which features a solid longitu- being 0,4 to 3,2 cm long and 1,1 to 1,4 cm wide. dinal nervure (Fig. 3b). From the widely decurrent These pines present an open venation, derived from (Fig. 3e), separated (Fig. 3f) to slightly imbricated the rachis nervures, which is denser at its base and (Fig. 3b) rachis, minor sessile pines displaying com- that disperses along the pinnules (Fig. 3a,b,c and plete limbus emanate with a somewhat acute angle d). As the nervures go along the pinnules they suf- of insertion (50˚ a 70˚). The pines outline is subcir- fer dichotomy, which can occur twice or four times cular and either smooth (Fig. 3f) or showing small (Fig. 3a). Pinnules are strongly decurrent and the insertions (Fig. 3a,b, and d). These insertions origi- more developed ones are free down to their base and

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may present an asymmetrical, subcircular to slightly The samples comprise basal and medial-basal oblong outline. fragments of elongated and bipinnate fronds (3,8 The associated microflora, composed of trilete to 9,0 cm long and 0,6 to 7,9 cm wide). They spores (75%) and monosaccate pollen (15%), rare present a robust main rachis, with strong longitu- bisaccate and striate pollens, was related by An- dinal nervures, which can be 0,9 cm wide at its dreis et al. (1979) to the H1/H2 biostratigraphic proximal portion and disperse at the more distal interval of Daemon and Quadros (1970). It cor- pines (Fig. 3b). In a subordinate way, and show- responds to the upper part of the Potonieisporites- ing somewhat acute insertion angles (50˚ to 70˚), Lundbladispora Zone proposed by Archangel- sessile, widely decurrent to pseudopetiolate pines sky and Marques-Toigo (1978) for the Chaco- emanate from the rachis. They change from iso- Paraná Basin (Argentina). According to Andreis et lated to slightly imbricated, show complete limbus, al. (1979), this interval is equivalent to either the and can either present or lack insertions on their bor- Stephanian C – Sakmarian boundary or, according ders, therefore producing their bipinnate aspect. to Faddeieva (1976), to the base of the Asselian in The pines from the basal portions of the fronds Russia. are decurrent and separated (Fig. 3e) to fairly im- The outcrop Base Morro Papaléo is situated in bricated (Fig. 3b). They can present rhomboidal the Barão do Triunfo topographic sheet (SH-22-0-1- (Fig. 3e), rhomboidal-ovoid (Fig. 3f) or rhomboidal- 2), 8 Km northwestward of Mariana Pimentel. It is elongated (Fig. 3b) outlines with no insertions. They located in a ravine on the lower part of the southwest- are 0,8–1,8 cm long and 1,3–1,8 cm wide. These ern border of the Morro Papaléo (UTM 0438317 E pines present an open venation, derived from the and 6647584 N). rachis nervures, which is denser at its base, disperses This outcrop, described by Pasqualini et al. at the apex and can dichotomize twice to three times. (1986), comprises an alternation of carbonaceous, The pines from the medial-basal portions of the fossiliferous siltstone, mica-rich, fine-grained sand- fronds are rather imbricated and change from exten- stone, lenticular, coarse-grained sandstone and con- sively decurrent to pseudopetiolate. They present ei- glomerate, and sandstone exhibiting iron-rich con- ther an elliptical-elongated or oblong-elliptical out- cretions. Plane bedded siltstone and sandstone and line that can display slight to strong insertions on cross-bedded sandstone are common sedimentary their borders. In this latter case a bipinnate mor- features. phology is produced and the lobes correspond to Pasqualini et al. (1986) related the fossiliferous the pinnules. The pine length spans from 0,7 to 5,4 beds to the bottom of efemerous water bodies asso- cm and their width between 0,9 and 2,8 cm. They ciated with the margins of interdistributary lakes. present an open venation, derived from the rachis The taphoflora is mostly composed of glos- nervures, which is denser at its base and more dis- sopterids with pinnate venation (Glossopteris com- perse along the pinnules. As the nervures go along munis, Glossopteris indica) and Botrychiopsis plan- the pinnules they suffer dichotomy, from twice to tiana associated with articulates (Phylloteca indica) four times. The pinnules are strongly decurrent and and conifers (Buriadia isophylla). Gangamopteris the more developed ones are free down to their base. sp., Glossopteris angustifolia, Cordaites and pro- They always present an asymmetrical outline that toglossopterids (Rubidgea obovata) occur as minor can be subcircular to slightly oblong (Fig. 3). elements. One of the samples (PB 2575), previously de- Some specimens formerly described by Pas- scribed by Guerra-Sommer et al. (1986), presents qualini et al. (1986) were analyzed herein, as well the central axis of the plant perpendicularly posi- as others stored in the Paleobotanic Sector of the tioned relative to the bedding planes and from which Earth Science Institute at UFRGS. four fronds radiate (3,3 cm long and 1,8 cm wide)

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Fig. 3 – Botrychiopsis plantiana fronds from Faxinal and Base of Morro Papaléo: (a) apical portion of the frond with pines whit complete limbus, presenting small insertions and venation - PB 2677; (b) detail from the average portion of a frond – PB 2935; (c) detail from pines and it insertion in the rachis – PB 3009; (d) detail of a pine with venation – PB 3097; (e) basal portion of the frond – PB 2679; (f) basal portion of the frond, representing the morphological differentiation – PB 2499.

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and a rachis (0,6 cm wide) with strong longitudinal The samples are stored in the Botany and Pa- nervures. From the rachis subaltern, separated, ses- leobotany Sector of the UNIVATES Natural Sci- sile, extensively decurrent pines emanate, exhibiting ences Museum and in the Paleobotany Sector of the entire limbus, slightly acute insertion angles (50˚ a DPE/UFRGS. 70˚) and a reniform morphology. These pines are The samples from Quitéria include basal and 0.6 cm long and 1.3 cm wide and show venation medial-basal fragments of elliptical, elongated and derived for the rachis nervures. Venation is open, bipinnate fronds (3,5 to 11,6 cm long and 0,4 to denser at its base (dispersing towards the apex) and 5,1 cm wide). They present a robust main rachis can dichotomize twice to three times. (0,1 to 0,9 cm thick) that encompasses strong lon- Samples from Base of Morro do Papaléo lack gitudinal nervures (Fig. 4 and Fig. 5a) from which spores and pollen probably due to non-preservation. sessile pines in a sub-opposite trend emanate with The Quitéria outcrop is located in the Pan- slightly acute insertion angles (60˚ to 80˚). These tano Grande County (Rio Grande do Sul State), in pines are decurrent to extensively decurrent, sepa- the southeastern border of the Paraná Basin (UTM rated to slightly imbricated, and present complete 0387516 E and 6643183 N). It is about 130 km from limbus, with (bipinnate morphology) or without in- Porto Alegre, along the BR 290 freeway, and can be sertions on their borders. localized in the Quitéria topographic sheet (SH22- The pines of the frond basal portions are widely NH). It comprises a sedimentary package about 8 decurrent and either separated (Fig. 4b and d and meters thick, characterized by the intercalation of Fig. 5c) or slightly imbricated (Fig. 4a and c and carbonaceous shale, argillite, fine- to coarse-grained Fig. 5a). They present a rhomboidal (Fig. 4d and sandstone, matrix-supported (silty to sandy) con- Fig. 5c), rhomboidal-elongated (Fig. 4c and Fig. 5a), glomerate and thin, up to 25 cm thick coal beds. rhomboidal-widened (Fig. 4a) or reniform outline The fronds are preserved in a light-yellow, fine- (Fig. 4b and Fig. 5b) on which insertions are some- grained sandstone bed (45 cm thick) that exhibits times observable (Fig. 4 and Fig. 5). Pines are 0,4 plane bedding and is situated immediately above a to 1,1 cm long and 0,8 to 2,5 cm wide. Pine ve- carbonaceous argillite bed. Both horizontal and in- nation, derived from the rachis nervures, is open, situ lycophyte stems (Brasilodendron) occur in as- denser towards its base and disperse at its apex, and sociation with this level, as well as Coniferophyta, dichotomize twice to four times (Fig. 4 and Fig. 6). Cordaites, Filicophyta and scattered fragments of The pines of the medial-basal portions of the glossopterids (Rubidgea sp. and Glossopteris sp.). fronds are slightly imbricated and vary from de- Piccoli et al. (1991) related the sedimentary current (Fig. 6b) to extensively decurrent (Fig. 6a). succession to a delta system dominated by low- They present an elliptical or elliptical-elongated out- energy fluvial processes, with well-developed allu- line and can present insertions (bipinnate morphol- vial plain facies and peat-forming swamps in the ogy) on their borders (Fig. 6d). Their lengths vary interdistributary lowlands. from 0,5 to 5,0 cm and their width from 0,7 to 2,8 cm. Based on specific features, mostly those re- These pines display an open venation, derived from lated to lycophyte specimens of the Brasilodendron the rachis nervures, which is denser at its basal and genus, and previous regional stratigraphic studies central portions and disperses along the pinnules. As (Menezes 1994, Chaves et al. 1994, Della Fávera et the nervures go along the pinnules they dichotomize al. 1994, Holz 1995, Lopes and Lavina 1995, Holz twice to four times (Fig. 4 and Fig. 6). Pinnules are 1997) Jasper and Guerra-Sommer (1999) suggested strongly decurrent and the more developed ones are a barrier-lagoon system (and associated washover free down to their base, presenting a subcircular to fans), similar to the model formulated by Reison elliptical-elongated outline. (1992). A palynological study on samples collected in

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Fig. 4 – Botrychiopsis valida fronds from Quitéria. (a) middle portion of a frond with pines, showing the venation of the rachis, the insertion of the pines in the rachis and is venation – PbU 0281; (b) middle portion of the frond with complete limbus presenting small insertions and the venation of the rachis and pines – PbU 0233; (c) basal portion of the frond with the insertion of the pines and venation – PbU 0252; (d) basal portion of the frond with the insertion of the pines and venation – PbU 0062. coalbeds of Quitéria outcrop has shown the pres- resented by the genera Lundbladispora, Vallati- ence of well preserved, abundant and diverse mi- sporites, Cristatisporites and Kraeuselisporites, are croflora. It is mainly composed of trilete spores, the dominant elements. Apiculati and Murornati 70% produced by pteridophytes, less frequent to rare spores were less common and point to the presence Gymnospermae pollens, alga, acritarchs and incer- of Filicophyta and Sphenophyta and are represented tae sedis. Zonati and Cingulicavati spores, related to by forms linked to the Punctatisporites, Calamo- herbaceous and shrub-like Lycophyta, mainly rep- spora, Leiotriletes, Deltoidospora, Murospora,

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Fig. 5 – Botrychiopsis valida.fronds from Quitéria (a) detail of the middle portion of a frond with pines and it insertion in the rachis – PbU 0281; (b) middle portion of the frond with complete limbus presenting small insertions and venation – PbU 0233; (c) basal portion of the frond with the insertion of the pines – PbU 0062; (d) apical portion of the frond with pines presenting a complete limbus with insertions and venation – PbU 0061.

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Fig. 6 – Details of pines of Botrychiopsis valida from Quitéria (a) detail of an apical pine with venation – PbU 0061; (b) detail of a basal pine – PbU 0062; (c) detail of a pine with venation – PbU 0233; (d) detail of venation of a basal pine, showing venation dichotomy – PbU 0062.

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Apiculatisporis, Granulatisporites, Cyclogranis- cimens indicate affinities with the Eusphenopteris porites and Convolutispora genera. Novik (1947) and Botrychiopsis (Kurtz) Archangel- Monosaccate pollen (Plicatipollenites, Cahe- sky and Arrondo (1971). niasaccites, Potonieisporites and Cannanoropollis The Eusphenopteris Novik (1947) genus is genera) is less common and indicates the occur- characterized by a bipinnate frond, with alate rence of Cordaitophyta. On the other hand, bisac- rachis and alternated pines. These pines show a cate pollen, such as Scheuringipollenites, Vesicas- poorly defined rachis as the pinnule limbus presents pora and Limitisporites, is related to Glossopterido- decurrent spreading. Pinnules are trilobate and phyta. deeply incised, with an open, dichotomized vena- The identified striated pollen specimens have a tion that arises from a central nervure starting from botanical affinity with Coniferophyta, such as Proto- the rachis. Within Carboniferous Gondwana succes- haploxypinus, Vittatina and Lunatisporites. Mono- sions these forms have been considered intermediate lete spores are rare and essentially represented by between Botrychiopsis and Dactylophylum (Zam- forms of Cycadopites. pirolli 2001). In Western Gondwana this genus was Due to the predominance of species related to recorded by Cúneo (1990), in the Mojón de Hierro Zonati and Cingulicavati spores the palynoflora is Formation (Upper Paleozoic Tepuel Group), Arroyo comparable, in the Paraná Basin, to the microfloris- Garrido area (Chubut Province ofArgentina). It was tic association identified in the Candiota Superior also found by Zampirolli (2001), in the Santa Marta and Banco Louco beds in the Candiota Coalfield by Farm (Itapeva region in the São Paulo State), within Meyer (1999). Neo-Carboniferous interglacial rocks (Itararé Sub- Among the spore-pollinnic associations pro- group of the Paraná Basin). posed by Marques-Toigo and Corrêa da Silva (1984) Based on previous studies (Archangelsky for the coal measures of the Rio Grande do Sul State, et al. 1987, Archangelsky and Cúneo 1991, this type of palynoflora is comparable to the Pun- Iannuzzi 1994, Ricardi 1994, Iannuzzi et al. 1998), tatisporites, Lundbladispora and Portalites Associ- Iannuzzi and Rösler (2000) have stated that the ation, which is related to plants of hydrophylous to Eusphenopteris genus was a rare component of the hygrophilous environments. Archeosigillaria/ Lepidondrops/ Frenguellia, Euro- In terms of biostratigraphy, the microflora american and Northern Rhacopteris Nothorha- identified in the Candiota Upper Coal Seams was copteris floras, although common in the Southern included in the Caheniasaccites ovatus Zone of Mar- Nothorhachopteris Flora, during the Carboniferous. ques-Toigo (1988) – Artinskian/Kungurian. Based Among the main features of the Eusphenopteris on isotopic dating of tonstein beds interbedded in genus are its tripinnate frond with trilobate pinnules, the coal seams, Cazzulo-Klepzig (2001) linked this deeply incised margins and subtriangular outline, zone to Late Kungurian/Early Roadian age. showing an open and dichotomized venation that Palynological analyses on samples from Quité- arises from a single and thick nervure going into ria confirm its correspondance with the Caheniasac- the pinnules from the rachis. Although presenting cites ovatus biozone (Kungurian/Early Roadian). an open and dichotomized venation in the pinnules, the studied material does not show all other features of the Eusphenopteris genus. TAXONOMIC AFFINITIES The morphology of the analyzed material In this chapter, the taxonomic affinities of the previ- points towards the Botrychiopsis (Kurtz) Archan- ously described material (Faxinal, base of the Morro gelsky and Arrondo (1971). Papaléo and Quitéria) are re-evaluated. A review of Paleozoic fronds related to Botry- The morphological features of the studied spe- chiopsis, taking into account not only morpho-

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logical and taxonomic data, but also stratigraphic by Archangelsky and Arrondo (1971), this species implications, was presented by Archangelsky and presents divided basal frond pinnules, confluent Arrondo (1971). The authors, based on Argentinean along the entire frond and presenting subrhombic material and relating it to Gondwana and extra- morphology with sinuous margins. These charac- Gondwana data, confirmed Kurtz’s (1895a, b) orig- teristics are not present in the studied samples. inal interpretations. A new diagnosis was defined The main morphological features of this mate- and the concepts of frond, pines and pinnules were rial are apical, medial and medial-basal fragments reviewed. Furthermore, descriptive features diag- of the pines. The pines have long and bipinnate nostic of the following three species were presented: fronds presenting a robust main rachis that can dis- Botrychiopsis weissiana (Kurtz) Archangelsky and perse in the distal pines. The rachis presents strong Arrondo (1971), Botrychiopsis plantiana (Carru- longitudinal nervures from which minor sessile pin- thers) Archangelsky and Arrondo (1971) and Botry- nules radiate displaying slightly acute insertion an- chiopsis valida (Feistmantel) Archangelsky and Ar- gles (50˚ to 70˚). The pinnules are generally decur- rondo (1971), the latter only compared with the Up- rent to pseudopetiolate, separated to slightly imbri- per Paleozoic of India. According to Archangelsky cated, presenting entire limbus with or without inser- and Arrondo (1971) the different species present a tions on their borders. They may assume a sublobate wide geographic distribution and clear chronostrati- morphology, from rhomboidal, rhomboidal-ovaled graphic boundaries that allow some phylogenetic as- or rhomboidal-elongated to elliptical, elliptical- sumptions in terms of an ascendancy of Botrychiop- elongated or oblong-elliptical morphology. They sis weissiana in relation to Botrychiopsis plantiana, present an open venation derived from the rachis and of this one to Botrychiopsis valida. nervures, which is denser at their base and disperse Cúneo (1990) confirms the genus features along their limbus, which can dichotomize twice pointing out that the pines are arranged in a slightly to four times. These elements link this material subopposite way, with a rachis that goes to the base to Botrychiopsis plantiana (Archangelsky and Ar- of the apical pinnule. Furthermore, pines are lobate rondo 1971). with deccurent insertions, its venation is open and This conclusion is reinforced by comparative can be dichotomized up to three times. analysis of the samples discussed herein and the In this study, both generic and specific propo- specimens shown by Archangelsky and Arrondo sitions of Archangelsky and Arrondo (1971) are ac- (1971, Slide V, Fig. 1 and 2, Slide VI, Fig. 1, 2 and cepted. A pteridophytic or pteridospermic affinity 3) and their diagnoses. has been proposed for this genus. This review also confirms the designations of Considering the original diagnosis of Kurtz Guerra-Sommer et al. (1980), Cazzulo-Klepzig and (1895a,b), emended by Archangelsky and Arrondo Guerra-Sommer (1983) and Pasqualini et al. (1986). (1971), Botrychiopsis weissiana presents some fea- Botrychiopsis valida tures that were not seen in the studied samples of the Faxinal and Base of Morro Papaléo outcrops. The The Quitéria frond samples were not correlated to morphological characters of Botrychiopsis weis- Botrychiopsis weissiana due to the lack of some crit- siana include an alate basal portion of the frond, a ical morphological features, such as the basal por- twisted insertion of the pines with a pronounced im- tion of the alate frond, the twisted insertion of the brication, and an ovoid-spatulated morphology of pines displaying heavily delineated imbrication and the frond apical pinnule. the ovoid-spatulated morphology of the frond apical Similarly, it is not possible to correlate the stud- pinnules. ied samples with Botrychiopsis valida. In Feist- On the other hand, the Quitéria forms can not mantel’s (1879) previous description, also emended be associated with Botrychiopsis plantiana due to

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the absence of characteristics such as the connected Botrychiopsis valida was previously described to slightly imbricate pinnules of the frond base and as Neuropteris valida by Feistmantel (1876) within the spatulate shape with the rounded distal margin Eopermian sedimentary successions of India. Later, of the apical pinnules. Feistmantel (1879, 1880) again mentioned the same However, these specimens can be correlated to species, also within Indian sedimentary rocks. another species, Botrychiopsis valida, taking into Gothan (1927) registered the presence of Gondwani- account the material described by Feistmantel dium validum in the Serra de los Llanos (Argentina). (1880), Archangelsky and Arrondo (1971), Rigby Archangelsky and Arrondo (1971) synonymized all (1973), Anderson and Anderson (1985) Gould these specimens to Botrychiopsis valida, stressing (1975), Retallack (1980), Rayner (1995), Rigby that this species was restricted to India. In their spe- (1985) and Archangelsky and Cúneo (1981). cific description, Archangelsky and Arrondo (1971) The pines and pinnules morphology along the highlight that the frond base presents well-defined, fronds supports the identification of the Quitéria ma- subcircular to slightly elongated and clearly sepa- terial with Botrychiopsis valida. Among the main rated pinnules with a large insertion base. The pines features used for its identification the presence of a present normal insertions and vary from slightly im- robust and well-defined main rachis can be consid- bricate to united. The highest number of pinnules ered, with solid longitudinal nervures, from which per pine (up to 5 cm long) is five. The usual in- subopposite sessile pinnules emanate with slightly sertion angles range from 70˚ to 80˚. All pinnules acute insertion angles (60˚ to 80˚). These pinnules are convergent, the apical ones presenting a sub- are extensively decurrent to decurrent, separated to rhombic morphology with sinuous margins. Ac- slightly imbricated, and have entire limbus present- cording to Archangelsky and Arrondo (1971), an- ing, or not, insertions on their borders. other important aspect of this species is the pres- Furthermore, the decurrent aspect of the sepa- ence in the medial portions of the frond of just pro- rated basal pines, with rhomboidal, elongate- duced pines. These pines present fused pinnules rhomboidal or reniform outline, are very similar at their base and free ones at their apical portions. to the material previously related to Botrychiopsis Similar features are reported by Anderson and An- plantiana by Gould (1975, Fig. 3C and D) and later derson (1985, pl. 167, Fig. 2) and can be seen in on to Botrychiopsis valida by Retallack (1980). the material presented herein. However, Retallack Some other aspects also evidence the associa- (1980) considered the designation of Botrychiop- tion of the specimen hereby analyzed to Botrychiop- sis valida, defended by Archangelsky and Arrondo sis valida, largely by comparison with the mate- (1971), as nomen vanum and sinonymized to Botry- rial described and synonymized by Archangelsky chiopsis ovata (Mc’Coy). Retallack (1980) also and Arrondo (1971) as well as that identified by states that the specimen classified as Botrychiop- Rigby (1973), Anderson and Anderson (1985), sis plantiana by Gould (1975, Fig. 3C, D) should Gould (1975), Retallack (1980) and Rayner (1995). also correspond to the Botrychiopsis ovata species. Among these evidences, the presence of slightly Later, Rigby (1985) suggests, keeping the Botry- imbricate, highly decurrent to decurrent pinnules chiopsis ovata species, including in this species with elliptical or elongated-elliptical outline and other taxa such as Otopteris ovata Mc’Coy (1847, sub-lobate border insertions was observed. The pin- p. 148; pl. IX, Fig. 2), Aneimites austrina Etheridge nules present an open venation, derived from the (1888, p. 1304; pl. 37), Cardiopteris polymorpha in rachis nervures, denser at their basal and central White (1969a,b; p. 98; platesA and B, Fig. 1), Gond- portions and dispersing along the upper portions of wanidium plantianum in Rigby (1973, p. 4-5; plate the pinnules, which dichotomize twice to four times 1, Fig. 2 and plate 2 Fig. 3), Triphyllopteris austrina (similar to the basal pinnules). in Morris (1975, p. 104; pl. 8.1c, Fig. 8.1 and 8.4

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k, u), Botrychiopsis in Retallack (1980, p. 394-395; chronostratigraphic impact Archangelsky and Ar- Fig. 21.3 E-G) and Otopteris ovata in Archangelsky rondo (1971) indicated a time span from Upper Car- (1983, p. 161-163; pl. 1-3). boniferous (Westphalian) to Lower Permian (Artin- Even so, in this paper the designation Botry- skian) for the genus. The Botrychiopsis weissiana chiopsis valida is maintained in accordance with biochron was restricted to the Carboniferous Sys- Archangelsky and Arrondo (1971) criteria. tem (Tupense local age – Westphalian/Stephanian of Western Europe). This geochronological posi- THE BIOSTRATIGRAPHIC IMPLICATIONS tion was corroborated by marine invertebrates (An- OF THE Botrychiopsis GENUS IN telo 1972) and palynology (Menéndez 1965). SOUTHERN PARANÁ BASIN Botrychiopsis plantiana is linked to the earliest The Botrychiopsis genus was proposed by Kurtz Conifers and Ginkgoales occurrences in the Tram- (1895a,b), based on Carboniferous material of the peadero and Libertad formations (Paganzo Basin), Rio Jejenes Formation (San Juan Province, Argen- in Upper Carboniferous sequences of Trampead- tina). Similar forms, previously recorded by Fren- erense local age. güelli (1944, 1946, 1954) under other denomina- The record of Botrychiopsis plantiana in the tions, were identified by Archangelsky and Arrondo Nueva Lubecka Formation, referred to the Lube- (1971) as Botrychiopsis weissiana. kenseA/B (Sakmarian/Artinskian), according to pa- According to Archangelsky and Arrondo lynological and marine invertebrates (Antelo 1972), (1971), the description of distinct biocrons for the is considered as a Carboniferous relict. species of the Botrychiopsis genus highlights not just Archangelsky and Arrondo (1971) considered its phylogenetic relationship but also its chronos- Botrychiopsis valida as exclusive to Indian succes- tratigraphic importance. The phylogenetic associ- sions (Kaharbari Formation). The age interval was ations come from the Botrychiopsis weissiana and attributed to the Artinskian through the analysis of Botrychiopsis plantiana sequential record within marine invertebrates (Robison 1967), hence com- Argentinean Gondwana associations. Forms related prising the youngest species of the genus (Archan- to Botrychiopsis valida would also be descended gelsky and Arrondo 1971). from Botrychiopsis weissiana developed in distant In the Paraná Basin, Carruthers (1869) regis- areas, hence differentiating from the Western Gond- tered Botrychiopsis plantiana in coal-bearing strata, wana coeval forms. Based on these evidences and later named Odontopteris plantiana. White (1908) the inferences of Gothan and Sahni (1937), it seems and Lundqvist (1919) designated the same form with that the group development occurred from an ances- the epithet Neuropteridium plantianun. Dolianiti tral stock identified in Peninsular India andAustralia (1948) named similar species found in the Itararé Namurian successions. Group of the Rio Grande do Sul State (Acampa- However, some aspects of this evidence, such mento Velho outcrop) as Gondwanidium plan- as the presence of Botrychiopsis weissiana and tianum. Botrychiopsis plantiana as coeval elements within In an attempt to characterize a Pre-Glossop- Argentinean Paleozoic floras (Archangelsky and teris Flora for Upper Paleozoic successions of the Azcuy 1985), suggest the need of re-evaluating pre- Northern Paraná Basin (Monte Mor outcrop – Itararé supposed phylogenetic relations. Although these Group, São Paulo State), Millan (1975, 1979) data support the idea that the specific relations pointed out Botrychiopsis plantiana as the diagnos- should be reviewed, studies on distinct basins have tic element. A Westphalian/Stephanian age was pro- confirmed the stratigraphic importance of the differ- posed for the succession based on palynological data ent morpho-species. (Trindade 1970). According to Iannuzzi (pess. In a first inference to the Botrychiopsis comm. 2002), however, from the analysis of the

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available type-material studied by Millan (1975, of the formerly described Santa Marta Taphoflora 1979), which is now kept in the Museu Nacional (Millan 1987a), using the same material that is now (UFRJ), it is impossible to establish the specific stored in the National Museum (Universidade Fed- affinity of the material. Therefore, the specimens de- eral do Rio de Janeiro). In this review, in addition to scribed by Millan (1975, 1979) remain here termed Botrychiopsis, other important diagnostic elements Botrychiopsis sp. have been included, such as Sphenopteris sp. and On the informal phytostratigraphic zonation Nothorhacopteris cf. argentinica. Palynological of the Late Paleozoic Paraná Basin, Rösler (1978) data from the same strata are comparable with the recorded the presence of Botrychiopsis plantiana Ahrensisporites cristatus Interval Zone of the Itararé in Taphoflora A, relating it to the early stages of Group lower portion (Zampirolli et al. 2000, Souza deglaciation in the Paraná Basin (Sakmarian/ 2000). Taking in account mega- and microfloristic Artinskian). Andreis and Archangelsky (1996), in a data, Zampirolli (2001) estimated a Westphalian age review of the Neopaleozoic basins of South Amer- for the Santa Marta Flora. According to Zampirolli ica, related the Taphoflora A to a probable Stepha- (2001) this association is comparable to the NBG nian age, taking into account the lack of the Glos- Zone of Argentina (Middle to Late Carboniferous sopteris genus. According to Guerra-Sommer and – Tupense – according to Archangelsky and Cúneo Cazzulo-Klepzig (1981) the Itararé Group tapho- 1991). flora in the Rio Grande do Sul State, including Botry- In a study on the biostratigraphic importance of chiopsis plantiana, can be related to the Cerquilho the upper Paleozoic taphofloras in Southern Paraná (Taphoflora A and B of Rösler 1978) and Argen- Basin, Guerra-Sommer and Cazzulo-Klepzig (1993) tinean Bajo de Veliz (Lubekense A) taphoflora. characterized some morphogenera as very impor- Studies of Millan (1987a) on the megafloris- tant in terms of dating and correlation. In their at- tic association of coal-bearing strata from Itapeva tempt to establish paleofloristic associations based (São Paulo State) – the Santa Marta Flora of the on observable boundaries of lithostratigraphic units, Itararé Group – record a large amount of Botrychiop- Botrychiopsis plantiana (recorded in the Faxinal and sis plantiana fronds and subordinate Sphenopsida Francisquinho outcrops by Andreis et al. 1979) was (Paracalamites, Sphenophyllum), Cordaitales (Cor- regarded as one of the main elements related to the daites) and platispermics seeds (Cordaicarpus). An interval of deposition of the Itararé sequences in the Itapevense local age was indicated for this associa- Rio Grande do Sul State (Sakmarian). tion. Botrychiopsis plantiana remains a significant In an attempt to define floristic stages for coal- component of the Eopermian Glossopteris flora, beds of the Itararé Group, Millan (1987b) correlated then associated with Glossopteris, Gangamopteris, the Monte Mor Taphoflora (Montemorense Stage) Phyllotheca, Buriadia (Pasqualini et al. 1986), to the Trampeaderense Megafloristic Zone of which was related by Guerra-Sommer et al. (1986) the Argentinean Paganzo Basin (Azcuy and Jelin to the basal sequences of the Rio Bonito Formation. 1980). This taphoflora is regarded as Westphalian/ In a preliminary biozoning of the South- Stephanian and correlated to the Potonieisporites- ern Paraná Basin (Guerra-Sommer and Cazzulo- Lundbladispora Zone (Archangelsky and Césari Klepzig 1993) the Botrychiopsis plantiana Zone was 1986). According to Millan (1987b), Botrychiopsis identified (Itararé Group and base of the Rio Bonito plantiana is the leading element of the Itapevense Formation) and related to the Sakmarian-Artinskian. Sub-Stage (Itapeva Flora). A Stephanian age for the The main feature of the lower subzone (Gang- Itapevense Sub-Stage was estimated by correlation amopteris angustifolia) is the predominance of with palynological data (Lima et al. 1976). Botrychiopsis plantiana associated with protoglos- Zampirolli (2001) carried out a detailed review sopterids (Rubidgea obovata and Rubidgea lancio-

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Fig. 7 – Palaeomaps of megafloristic associations in biomes at cold-cool, cool (I-II) and warm temperate (III) climates in the Lower and Midle/Upper Permian (Adapted from Scotese 2002). latus) and the first glossopterids, mostly represented quence was closely related to palaeoclimate, in addi- by Gangamopteris (Gangamopteris obovata, Gang- tion to biostratigraphic and paleoecological controls amopteris buriadica and Gangamopteris angusti- (Guerra-Sommer et al. 2001). This assumption was folia). Glossopteris (Glossopteris communis and confirmed by the evaluation of taphofloristic param- Glossopteris indica) occur as complementary ele- eters from different biostratigraphic units. ments. The Phyllotheca indica Subzone comprises The homogeneous composition of Early Per- elements related to articulate (Phyllotheca indica) mian (Sakmarian) wet lowland biomes, character- associated with Glossopteris (Glossopteris commu- ized by the Glossopteris flora and herbaceous to nis) and Conifers (Buriadia isophylla). shrub-like plants considered to be relicts from a rig- Recent data have shown that paleofloristic evo- orous cold climate (e.g. Botrychiopsis plantiana), lution in the Southern Paraná Basin during deposi- suggests the persistence of an ice age. The progres- tion of the Carboniferous – Early Triassic Megase- sive colonization of the lowland clastic habitats by

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Fig. 8 – Megafloristic associations in biomes at cold-cool, cool and warm temperate climates in the Lower and Midle/Upper Permian in Southernmost Paraná Basin, Brazil. (1) Predominant; (2) Abundant; (3) Commom; (4) Less commom; (5) Rare. (I) Asselian wet biome; (II) Sakmarian/Artinskian wet biome; (III) Kungurian wet biome.

Cordaitales, Ginkgoales, Coniferales, in addition to cladus?) and arborescent Lycophyta (Brasiloden- the herbaceous articulates, indicates an Early Artin- dron pedroanum), as presented herein, indicates a skian climate warming. larger biogeographic distribution range of this genus Glossopterids with pinnate venation and re- in the Gondwana. lated to Glossopteris are common in Early Per- mian coal-bearing strata (Artinskian/Kungurian) CONCLUSIONS whereas Gangamopteris (palmate form) is poorly represented. The sudden enrichment of filicoid Botrychiopsis has been considered an impor- fronds is characteristic of the landscape units in this tant floristic element of Westphalian/Artinskian as- stage. Arborescent lycophyte communities become sociations of the Paraná Basin. In the Southern important, associated with glossopterid-dominated Paraná Basin, several authors have related its oc- communities. This event coincides with the wan- currence to glacial-influenced environments. This ing of the Permian icehouse stage, which reached genus has been recorded within associations related its peak around the Asselian/Sakmarian boundary. to a Permian glacier retreat (icehouse stage – As- The record of Botrychiopsis valida within coal- selian/Kungurian). During the early deglaciation bearing megafloras in the Southern Paraná Basin, phases (cold/cool temperate climate), the Glos- associated with Glossopterids (Glossopteris, Gang- sopteris Flora, linked to wet biomes, changed gradu- amopteris and Rubidgea), Filicophyta (Rhodeop- ally. However, during the latest deglaciation stages, teridium?), Conifers (Buriadia and Ferruglio- when a fast transition took place in the wet biomes

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Fig. 9 – Botrychiopsis Zone and related plant-assemblages, palaeoclimate, sequence stratigraphy units, relative sea level and litostrati- graphic units of Early Permian in Southern Paraná Basin, Brazil. due to warming (warm temperate climate), in asso- Zone encompassing the Asselian/Artinskian Botry- ciation with peat formation, the Botrychiopsis genus chiopsis plantiana and the upper Artinskian/ persisted, then represented by Botrychiopsis valida. Kungurian Botrychiopsis valida subzones (Fig. 9). In the phytostratigraphie scheme presented by The Faxinal microflora associated with the Guerra-Sommer and Cazzulo-Klepzig (1993), the Botrychiopsis plantiana Subzone suggests a Stepha- Botrychiopsis plantiana Zone ranges from Sakmar- nian C/Sakmarian or lower Asselian age. The Qui- ian to Artinskian. téria microflora on the other hand, is related to the The new data here presented enlarge the Botry- Caheniasaccites ovatus Zone (Kungurian to lower chiopsis genus biochron and define a stratigraphic Roadian). hierarchy between Botrychiopsis plantiana and Botrychiopsis valida. In the Southern Paraná Basin, ACKNOWLEDGMENTS Botrychiopsis plantiana is restricted to Asselian/ André Jasper received support from UNIVATES, Artinskian and Botrychiopsis valida to Kungurian. FUNADESP, FAPERGS and CNPq, and Margot It is also reported that the stratigraphic range of the Guerra-Sommer, Miriam Cazzulo-Klepzig and Ru- Botrychiopsis genus in Southern Paraná Basin is re- aldo Menegat from FAPERGS and CNPq. lated to a deglaciation stage (icehouse) and, there- fore, to a wide climate range, from cold/cool tem- RESUMO perate to warm temperate. Megafloristic association O gênero Botrychiopsis tem sido considerado um ele- in biomes cold-cool, cool and warm-temperate cli- mento florístico importante das associações do intervalo mates were identified, in the Lower and Midle/ Westphaliano/Artinskiano da Bacia do Paraná. O registro Upper Permian in Southernwest Paraná Basin (Fig. 7 de formas relacionadas ao gênero Botrychiopsis, especifi- and 8). camente Botrychiopsis valida,emroof-shales na área de A new phytostratigraphic scheme is proposed. Quitéria, Formação Rio Bonito, no sul da Bacia do Paraná It includes an Asselian/Artinskian Botrychiopsis amplia o biocron do gênero, definindo uma hierarquia

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estratigráfica para as espécies Botrychiopsis plantiana e Archangelsky S. 1984. Floras neopaleozoicas del Botrychiopsis valida para esta bacia. A distribuição es- Gondwana y su zonación estratigráfica: aspectos pa- tratigráfica do gênero está condicionada a controle climá- leogeográficos conexos: In: Lemos de Souza MJ. tico relacionado a um ciclo de deglaciação em estágio ice- (Ed), Proceedings and Papers of the Symposium on Gondwana Coals, Lisbon, 1983: Comunicações dos house, com espectro de tolerância climática abrangente, Serviços Geológicos de Portugal, v. 70, p. 135-150. desde condições climáticas do tipo frio até temperado/ Archangelsky S and Arrondo OG. quente. É proposto um novo zoneamento fitoestratigrá- 1971. Paleo- phytologia Kurtziana III. 2. Estudio sobre el género fico para essa porção da bacia, incluindo uma Zona Botry- Botrychiopsis Kurtz (= Gondwanidium Gothan) del chiopsis (Asseliano/Kunguriano) com duas sub-zonas Carbónico y Pérmico gondwánico. Ameghiniana 8: Botrychiopsis plantiana (Asseliano/Artinskiano) e Botry- 189-227. chiopsis valida (topo do Artinskiano/Kunguriano). Archangelsky S and Azcuy CL. 1985. Carboniferous Palavras-chave: Botrychiopsis, biostratigrafia, Bacia do paleobotany and palynology in Argentina. In: Car- Paraná, Permiano, paleoclimatologia, Gondwana. boniferous International Congress, 10. Pro- ceedings... Madrid, 1983 4: 267-280. Archangelsky S and Césari C. 1986. Comparación REFERENCES de palinofloras carboníferas de las Cuencas Paganzo Anderson JM and Anderson HM. 1985. Paleoflora (Argentina) y Paraná (Brasil). Bol Inst Geociências, of Southern Africa. Prodromus of South African Univ de São Paulo 17: 5-9. Megafloras, Devonian to Lower Cretaceous: A.A. Archangelsky S and Cúneo NR. 1981. Sobre la pres- Balkema, Rotterdam, 423 p. encia del genero Botrychiopsis Kurtz en la Forma- Andreis RR and Archangelsky S. 1996. The Neo- cion Nueva Lubecka, Permiano Inferior de Chubut, Paleozoic basins of southern south America. In: Argentina. In: Congresso Latino-Americano de Moullade M and Nairn AEM. The Phanerozoic Paleontologia, 2. Anais... Porto Alegre, p. 157- Geology of the World (Ed.), Amsterdam 5: 341-575. 167. Andreis RR, Cazzulo-Klepzig M, Guerra-Sommer Archangelsky S and Cúneo NR. 1987. Feruglio- M and Marques-Toigo M. 1979. Interpretação pa- cladeaceae, a new Conifer family from the Permian leoambiental e estudo paleobotânico e palinológico of Gondwana. Review of Palaeobotany and Palynol- do Grupo Itararé na área do Faxinal, município de ogy 51: 3-30. Guaíba, RS. In: Simpósio de Geologia do Nor- Archangelsky S and Cúneo NR. 1991. The Neopa- deste, 9, Resumos... p. 30. leozoic floristic succession from Northwestern Ar- Antelo B. 1972. Los braquiópodos del Carbonífero gentina. A new perspective. In: Ulbrich H and Superior de la Quebrada Larga, en las cabeceras del Rocha Campos AC. (Eds), Gondwana Seven, Pro- Rio Blanco, Provincia de San Juan. Ameghiniana 9: ceed. Instituto de Geociências, Universidade de São 159-172. Paulo, São Paulo, p. 469-481. Archangelsky S. 1971. Las tafofloras del Sistema Archangelsky S and Marques-Toigo M. 1978. La Paganzo en la Republica Argentina. An Acad Bras Palinologia y el problema del limite Carbonico- Cienc 48: 67-88. Permico en el Gondwana Sulamericano. In: Con- gresso Argentino de Paleontologia y Bioes- Archangelsky S. 1978. Paleoecologia del Paleozoico tratigrafia,2,yCongresso Latino-Americano Superior Argentino sobre la base de sus plantas fos- de Paleontologia, 1. Actas... Buenos Aires 1978, iles. Ameghiniana 15: 73-84. 4: 68-74. Archangelsky S. 1983. Nothorhacopteris, a new Archangelsky S, Azcuy CL, Gonçalvez CR and generic name for some Carboniferous monopinnate Sabattinin N. 1987. Paleontologia, Biostratigrafia fronds of Gondwanaland (= Rhacopteris ovata auct. y Paleologia de las cuencas Paganzo, Callingasta Us- and Pseudorhacopteris Rigby 1973). Palaeobot Pa- fallata y Rio Blanco. In: El Sistem Carbonífero lynol 38: 157-172.

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